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December 1997 Volume 15, N° 3

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Source : MNHN, Paris

AIVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

December 1997 Volume 15, N° 3

Alytes, 1997, 15 (3): 105-112. 105

À new species of Physalaemus
(Anura, Leptodactylidae)
from the Atlantic rain forest
of northeastern Brazil

José P. POMBAL, Jr. & Cristiane A. MADUREIRA

Departamento de Vertebrados, Museu Nacional, Quinta da Boa Vista,
20940-040 Rio de Janeiro, RJ, Brasil

A new species of leptodactylid frog is described from Passo de Cama-
ragibe, State of Alagoas, northeastern Brazil. The new species is a member
of the Physalaemus signifer group, and is characterized by its medium
size, snout protruding in lateral view, dorsolateral light fold extending from
the posterior corner of the eye to the inguinal region, and large outer

metatarsal tubercle. Description of the tadpole is provided. Bis] iothèque Centrale Muséum

3 185

3001 000503!

The genus Physalaemus is known from Mexico to southern South America, with 37
recognized species (FROST, 1985; DUELLMAN, 1993). Four species groups are currently rec-
ognized in the genus: P biligonigerus group, P cuvieri group, P. pustulosus group and P signifer
group (LyYNCH, 1970); however, P deimaticus Sazima & Caramaschi, 1986 and P rupestris
Caramaschi, Carcerelli & Feio, 1991 are not presently placed in any species group (SAZIMA &
CARAMASCHI, 1986; CARAMASCHI et al., 1991). The P signifer group is characterized by its
small to moderate size (15-35 mm SVL), slender body, smooth skin, first finger shorter than
the second, no inner tarsal tubercle, small, non-compressed metatarsal tubercles, small to
large inguinal glands, and parotoid glands absent (LYNCH, 1970). The species presently
allocated to the P signifer group are P. bokermanni Cardoso & Haddad, 1985, P. crombiei
Heyer & Wolf, 1989, P maculiventris (A. Lutz, 1925), P. moreirae (Miranda-Ribeiro, 1937), P.

_XK A
BIBL. DU
MUSÉUM
PARIS
* Source : MNHN, Paris

INTRODUCTION 3

106 ALYTES 15 (3)

nanus (Boulenger, 1888), P. obtectus Bokermann, 1966, P. olfersii (Lichtenstein & Martens,
1856), P signifer (Girard, 1853) (FROST, 1985; DUELLMAN, 1993), and P spiniger (Miranda-
Ribeiro, 1926) (see HADDAD & POMBAL, in press). Physalaemus franciscae Heyer, 1985 is a
synonym of P moreirae (CARAMASCHI & CARAMASCHI, 1991). Herein, we describe a new
species of the P signifer group from northeastern Brazil.

MATERIAL AND METHODS

Specimens used in the description or examined for comparisons are deposited in AL-MN
(Adolpho Lutz Collection, deposited in Museu Nacional, Rio de Janeiro, Brazil), CFBH
(Célio F. B. Haddad Collection, deposited in Departamento de Zoologia, Universidade
Estadual Paulista, Rio Claro, Brazil), MNRJ (Museu Nacional, Rio de Janeiro, Brazil),
ZUEC (Museu de Histéria Natural, Universidade Estadual de Campinas, Brazil). Additional
specimens examined are listed in app. 1.

Abbreviations used in the measurements of the adults are SVL (snout-vent length), HL
(head length), HW (head width), ED (eye diameter), IOD (interorbital distance), THL (thigh
length), TBL (tibia length), and FL (foot length). All measurements are in millimeters. The
measurements of the adults followed DUELLMAN (1970) and Cet (1980). Measured adult
specimens were fixed in 10 % formalin and preserved in 70 % ethyl alcohol. The tadpoles were
preserved in 5% formalin. For measurements we used an ocular micrometer in a Zeiss
stereomicroscope, except for SVL that was measured with a caliper. Drawings of the holotype
and tadpole were made using a Zeiss stereomicroscope with a drawing tube. Tooth row
formula of tadpoles is given according to ALTIG (1970).

RESULTS

Physalaemus caete sp. nov.

Holotype.- MNRIJ 9803, adult male (fig. 1), collected at Fazenda Santa Justina, Municipality
of Passo de Camaragibe (approximately 9°13'S 35°31°W; 45-90 m elevation), State of Ala-
goas, Brazil, on 26-30 June 1988 by Dante M. TEIXEIRA.

Paratopotypes. - MNRIJ 9801-02, 9804-05, 9848-50, adult males, collected with the holotype.

Diagnosis. - À medium-sized species (males 23.3-25.8 mm SVL) belonging to the Physalae-
mus signifer group (sensu LYNCH, 1970), characterized by: (1) short and wide head; (2) snout
nearly rounded in dorsal view and protruding in lateral view; (3) canthus rostralis weakly
distinct; (4) dorsolateral light fold extending from the posterior corner of the eye to the
inguinal region; (5) two or three black spots on the tibia; (6) large outer metatarsal tubercle.

Physalaemus caete is distinguished from P offersii by its smaller size (P. olfersii 28.5-
34.5 mm SVL; HEyer et al., 1990) and by the presence of dark inverted V-shaped marks on the
back (absent in P olfersii). By its larger size, the new species differs from P. bokermanni, P.
crombiei, P. maculiventris, P. nanus, P. signifer and P. spiniger (combined SVL ranging
15.0-22.3 mm; BOKERMANN, 1962; CARDOSO & HADDAD, 1985; HEYER & WOLF, 1989; HEYER
et al., 1990; HADDAD & POMBAL, in press; personal observations). Further, the new species
differs from P crombiei, P. maculiventris and P. signifer by its broader head. From P

Source : MNHN, Paris

POMBAL & MADUREIRA 107

Fig. 1. Physalaemus caete, holotype MNRJ 9803. (a) Dorsal view (b) ventral view.

bokermanni and P signifer, P. caete also differs in having a snout protruding in lateral view (see
P signifer and P. bokermanni figures in BOKERMANN, 1962 and CARDOsO & HADDAD, 1985,
respectively). Physalaemus caete is distinguished from P maculiventris and P. moreirae by its
relatively uniformly colored belly (boldly dark and light mottled posterior belly in P moreirae
and P maculiventris;, HEYER, 1985; personal observations). From P obtectus, P. caete is
distinguished by its larger outer metatarsal tubercle and its smaller head (in P obtectus head
length 31-32 % SVL, in P caete head length 22-24 % SVL).

Description of holotype. - Body nearly slender; head wider than long; snout nearly rounded in
dorsal view, protruding in lateral view (fig. 2a-b); nostrils slightly protuberant, directed
laterally; canthus rostralis weakly distinct; eye slightly protuberant; tympanum weakly dis-
tinct, large; distinct supratympanic fold from tympanum to shoulder; narrow dorsolateral
fold extending from the posterior corner of the eye to the inguinal region; vocal sac distinct,
subgular, slightly expanded externally, extending to the border of chest with belly; vocal slits
present; choanae large, nearly round; tongue narrow, long; vomerine teeth absent; maxillary
teeth not visible, but discernible by probe. Arms slender, forearms moderately robust; fingers
short, no prepollex; brown nuptial pad on each thumb; subarticular tubercles single, protrud-

Source : MNHN, Paris

108 ALYTES 15 (3)

Fig. 2. Physalaemus caete, holotype, MNRJ 9803. (a) Dorsal and (b) lateral views of head: ventral views
of (€) hand and (d) foot (scale = 2 mm).

Source : MNHN, Paris

PoMBAL & MADUREIRA 109

ing and round; outer metacarpal tubercle large, ovoid; inner metacarpal tubercle large, nearly
elliptical; small supranumerary tubercles; finger tips slightly expanded; finger lengths I = IV <
I < II (fig. 2c). Legs moderately robust; tibia length longer than thigh length; foot with a
protruding, elliptical inner metatarsal tubercle; outer metatarsal tubercle small, protruding
and round; well-developed subarticular tubercles, single, protruding and round; small supra-
numerary tubercles; toe tips slightly expanded; toe lengths I < II < V < III < IV (fig. 2d).
Inguinal glands small; dorsum skin and venter smooth.

Color of holotype. — In preservative, dorsum brown with three dark brown inverse V-shaped
marks with a fine light border, and also with dark brown dots with a fine light border; a white
line on the dorsolateral fold; flanks below the dorsolateral fold dark brown; inguinal gland
with a black spot on its right side and three black spots on the left side; forearm light brown
with a dark brown transverse bar; a dark brown spot below the elbow; thigh, tibia and foot
light brown; thigh and tibia with a dark brown transverse bar, foot with a dark brown spot;
black marks on the knee and upperparts of thigh; three black spots on the tibia and tarsus;
posterior tarsus and anal region black; sole of foot gray; palm cream; throat dark gray; belly
gray with small light gray dots.

In life, the inguinal region was orange-reddish (D. M. TÆIXEIRA, personal communica-
tion).

Measurements of the holotype. -SVL 24.5; HL 5.3; HW 8.7; ED 2.8; IOD 4.2; THL 11.2; TBL
12.3; FL 11.7.

Variation. — In preservative, dorsum brown to dark brown; in some specimens the dorsal
pattern is less evident or without the fine light borders of the inverse V-shaped marks; in one
individual the dorsum is rugose; inguinal glands well or poorly developed; the black spots on
the inguinal glands are distinct in all individuals, but sometimes are small. Measurements
(mean + standard deviation, range) of seven males (females unknown) are as follows: SVL
24.54 + 0.85, 23.5-25.8; HL 5.74 + 0.40, 5.3-6.2; HW 9.10 + 0.44, 8.5-9.6; ED 2.67 + 0.17,
2.4-2.0; IOD 4.26 + 0.20,4.0-4.6; THL 11.93 + 0.64, 11.2-12.7; TBL 12.31 + 0.21, 12.0-12.6;
FL 11.64 + 0.41, 11.0-12.2.

Tadpoles. — À foam nest where males of P caete were in calling activity was found in the type
locality. Larvae were obtained by development of the eggs of this nest (D. M. TEIXEIRA,
personal communication). The following description is based on a tadpole in developmental
stage 28 (GOsNER, 1960). Body depressed-globular in lateral view (fig. 3a), ovoid in dorsal and
ventral views (fig. 3b-c); body wider than high; snout rounded; eyes small, dorsolateral;
nostrils dorsal, small and rounded, about midway between the eyes and the tip of snout;
spiracle sinistral, its opening past middle of body; cloacal tube median, medium sized; caudal
musculature slender; dorsal fin originating on body; ventral fin wider than dorsal. Oral disc
(fig. 3d) directed ventrally and bordered by papillae, with a large gap on the anterior labium;
tooth row formula 2(2)/3(1); jaw sheath strongly developed and serrate; posterior sheath
V-shaped. In preservative, body gray; caudal musculature with scattered melanophores,
concentrated on the upper first third; fins translucent with scattered melanophores.

Measurements: total length 14.9; body length 8.9; body heigth 2.4; body width 3.7;
internarial distance 0.4; interorbital distance 1.0; eye-nostril distance 0.5; eye diameter 0.5.

Source : MNHN, Paris

110 ALYTES 15 (3)

Sins
SERRE van S

Fig. 3. - Tadpole of Physalaemus caete, stage 28; (a) lateral, (b) dorsal and (c) ventral views (scale = 2
mm); (d) oral disc (scale = 0. 5 mm).

Distribution. — The new species is known only from the type locality, and from the municipal-
ity of Murici (approximately 9°47’S 36°50°W; 650 m elevation), both in the State of Alagoas,
northeastern of Brazil. The other species of the Physalaemus signifer group are known from
the States of Rio Grande do Sul to Espirito Santo (FROST, 1985; DUELLMAN, 1993). The new
species represents the northernmost record for the P signifer group.

Source : MNHN, Paris

POMBAL & MADUREIRA 111

Natural history. — Physalaemus caete was collected by day on the leaf litter around ponds in
the Atlantic Rain Forest. The eggs, in foam nests, were observed in the water of ponds and in
the water of tree holes, near the ground.

Etymology.- Caetéis a Tupi indigenous name, here used as a noun in apposition. The specific
name is an allusion for the forest habitat (cad, forest; eté, true), where P caete and most of the
species of the P signifer group are collected. Caeté was also the name of the extinct Indians
from the region of the type locality.

RÉSUMÉ

Une nouvelle espèce de grenouille leptodactyle est décrite de Passo de Camaragibe,
commune de l’état de Alagoas, nord-est du Brésil. La nouvelle espèce, qui appartient au
groupe de Physalaemus signifer, est caractérisée par sa taille moyenne, son museau proémi-
nent en vue latérale, ses plis dorsolatéraux clairs s'étendant de l’extrémité postérieure de l’oeil
jusqu’à la région inguinale, et son tubercule métatarsien externe développé. Une description
du têtard est fournie.

ACKNOWLEDGMENTS

Dante M. TrIxEIRA collected the new species and provided valuable information; U. CARAMASCHI
and C. F. B. HADDAD made helpful suggestions on the manuscript; P. R. NASCIMENTO made the line
drawings; U. CARAMASCHI made the photos of the holotype; H. ZAHER translated the résumé; C. F. B.
HaDpaD and I. SazimA loaned and/or permitted study of material under their care.

LITERATURE CITED

ALTIG, R., 1970. — A key to the tadpoles of the continental United States and Canada. Herpetologica, 26:
180-207.
BOKERMANN, W. C. A., 1962. — Notas sôbre três espécies de Physalaemus (Amphibia, Salientia, Lepto-
dactylidae). Anais Acad. bras. Cien., 63-569.
CarAMASCHI, U. & CARAMASCHI, E. P., 1991. — Reassessment of the type-locality and synonymy of
Physalaemus moreirae (Miranda-Ribeiro, 1937) (Anura: Leptodactylidae). J Herp., 25: 107-108.
CARAMASCHI, U., CARCERELLI, L. C. & FE10, R. N., 1991. — A new species of Physalaemus (Anura:
Leptodactylidae) from Minas Gerais, Southeastern Brazil. Herpetologica, 47: 148-151.
CarDoso, À. J. & HaDDan, C. F. B., 1985. - Nova espécie de Physalaemus do grupo signiferus (Amphibia,
Anura, Leptodactylidae). Rev. brasil. Biol., 45: 33-37.
Ce, J. M., 1980. - Amphibians of Argentina. Monit. zoo. ital., (n.s.), Mon. 2: 1-609.
DuELLMAN, W. E., 1970. - The hylid frogs of Middle America. Mon. Mus. nat. Hist. Univ. Kansas,
+ 1-753, pl. 1-72.
1993. - Amphibian species of the world: additions and corrections. Univ. Kansas Mus. nat. Hist.
special Publ., 21: [i-ï] + i-ii + 1-372.
FRosr, D. R. (ed.), 1985. - Amphibian species of the world. Lawrence, Allen Press & Ass. Syst. Coll. [i-iv]
+i-v + 1-732.
Goswer, K. L., 1960. — A simplified table for staging anuran embryos and larvae with notes on
identification. Herpetologica, 16: 183-190.

Source : MNHN, Paris

112 ALYTES 15 (3)

Happan, C. F. B. & PoMBAL, J. P, Jr. in press. - Redescription of Physalaemus spiniger (Anura:
Leptodactylidae) and description of two new reproductive modes. J Herp. in press.

HEYER, W. R., 1985. - New species of frogs from Boracéia, Sao Paulo, Brazil. Proc. biol. Soc. Wash., 98:
657-671.

Heyer, W. R. & WoLr, A. J., 1989. — Physalaemus crombiei (Amphibia: Leptodactylidae), a new frog
species from Espirito Santo, Brazil with comments on the P signifer group. Proc. biol. Soc. Wash.,
102: 500-506.

HEYER, W. R., RAND, A. S., GONÇALVES DA CRUZ, C. A., PEIXOTO, O. L. & NELSON, C. E., 1990. — Frogs
of Boracëia. Arg. Zool., 31 (4): 231-410.

LyncH, J D., 1970. - Systematic status of the American leptodactylid frog genera Engystomops,
Eupemphix, and Physalaemus. Copeia, 1970: 488-496.

SAziMA, L. & CARAMASCHI, U., 1986. — Descriçäo de Physalaemus deimaticus, sp. n., e observaçôes sobre
comportamento deimätico em P nattereri (Steindn.) - Anura, Leptodactylidae. Rev. Biol., 13:
91-101.

APPENDIX 1

ADDITIONAL SPECIMENS EXAMINED

Physalaemus caete.- MNRJ 9712-17 (Murici, AL); MNRJ 18280 (lot of tadpoles, Passo de Camaragibe,

Physalaemus bokermanni. - ZUEC 4520-21 (paratypes, Santo André, SP).

Physalaemus crombiei, - MNRJ 17694-745 (Aracruz, ES).

Physalaemus maculiventris. — AL-MN 684 (syntype, “Alto da Serra de Cubatäo”); MNRJ 4228-30
(Paranapiacaba, SP); MNRJ 1797, 9975-10020 (Serra de Araraquara, PR).

Physalaemus moreirae. - MNRIJ 464 (holotype, Paranapiacaba, Santos, SP).

Physalaemus nanus. — CFBH 081 (Sao José, SC); MNRJ 12827-32 (Florianépolis, SC).

Physalaemus obtectus. = MNRJ 4025, 14206-07 (paratypes, Linhares, ES).

Physalaemus olfersii. — MNRJ 2428 (Parati, RJ); MNRJ 0482, 5525-26, 12826 (Teresopolis, RJ).

Physalaemus signifer. - MNRJ 1123, 6616-35 (Duque de Caxias, RJ); MNRJ 2753, 12461-62 (Barro
Branco, Duque de Caxias, RJ); MNRJ 2766, 12477-80 (Pendotiba, Niterôi, RJ); MNRJ 12837-42
(topotypes, Jacarepaguä, Rio de Janeiro, RJ).

Physalaemus spiniger. - CFBH 312-17, 319-21, MNRJ 18474, ZUEC 6878, 6881-82 (Cananéia, SP);
ZUEC 9333-43 (Caraguatatuba, SP); CFBH 2479 (Eldorado, SP); MNRJ 18475-76 (Guaraque-
çaba, PR); CFBH 307-10, 410, 835, MNRJ 18470-72, 18473, ZUEC 6876-77 (topotypes, Iguape,
SP); ZUEC 3250 (Jacupiranga, SP).

Corresponding editor: W. Ronald HEYER.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 15 (3): 113-120. 113

Un nuevo Leptodactylus
(Anura, Leptodactylidae)
de un bosque nublado
del oeste de Venezuela

Abraham MuARES-URRUTIA

Centro de Investigaciones en Ecologia y Zonas Aridas (CIEZA),
Universidad Francisco de Miranda, Apartado 7506, Coro 4101-A, Venezuela

A new species of the frog genus Leptodactylus is described from Cerro
Socopé, Municipio Mauroa, State of Falcôn, western Venezuela. This
species is tentatively included in the L. podicipinus-wagneri complex of
species. It can be distinguished from other members of this complex in
lacking dorsolateral folds, and in having males with subovoid snout tip,
females with snout tip rounded to almost truncated, and light posterior lip
stripe visible and well defined.

INTRODUCCIÉN

Las ranas silbadoras del género Leptodactylus estân ricamente representadas y amplia-
mente distribuidas en Venezuela, con 18 especies registradas por LA MaRCA (1992). Luego de
la revisiôn de HEYER (1994) del complejo de especies de L. podicipinus-wagneri, se eleva el
nümero de especies del género a 26. En este ültimo trabajo, el autor examiné numerosos
ejemplares asignables a este complejo de especies, muchos de ellos provenientes de una
amplia lista de localidades en Venezuela (la mayoria a elevaciones entre 850 y 1900 m de
altitud), entre las cuales se menciona al Estado Falcén, sin señalar una localidad especifica, y
comenta que “Within Venezuela, there are clusters of specimens from three small geographic
areas that appear to represent distinct OTUs” (HEYER, 1994: 46). Uno de estos ejemplares
proviene de la siguiente localidad: “Falcén: Carora, 84 km NW of, Cerro Socopé, 1260 m,
USNM 216804”.

Durante un trabajo de campo realizado por el autor en el Cerro Socopé, Estado Falcén, se
encontraron siete ejemplares de Leptodactylus sin pliegues dorsolaterales evidentes, y los
machos con un par de espinas queratinizadas en el pulgar de cada mano. Una vezcomparados
con las reseñas publicadas por HEYER (1994) y con ejemplares provenientes de Mérida, Andes
de Venezuela, y las tierras bajas de la regiôn del sur del Lago de Maracaibo, se evidencié que
estos ejemplares representan una especie aun no descrita.

Source : MNHN, Paris

114 ALYTES 15 (3)

MATERIALES Y MÉTODOS

La terminologia y las medidas son una combinacién a conveniencia de las propuestas de
RIVERO (1961) y HEYER (1994). El registro de la formula de membranacién pedal sigue la
metodologia sugerida por LA MARCA (1994). Las medidas se presentan para todos los
ejemplares en la tab. 1. Los criterios para determinar la condiciôn adulta y la clase de tamaño
de los ejemplares son los mismos empleados por HEYER (1994). Las siguientes abreviaciones
son utilizadas para designar las colecciones: AMU, nümeros de campo de Abraham MHARES-
URRUTIA; CIEZAH, Colecciôn Herpetolégica del Centro de Investigaciones en Ecologia y
Zonas Âridas, Universidad Francisco de Miranda, Venezuela; EBRG, Estacién Biolôgica de
Rancho Grande, Maracay, Venezuela; ULABG, Coleccion de Anfibios y Reptiles del Labo-
ratorio de Biogeografia, Universidad de Los Andes, Mérida, Venezuela; USNM, National
Museum of Natural History, Washington, Estados Unidos.

RESULTADOS

Leptodactylus magistris sp. nov.
(fig. 1)

Holotipo. - EBRG 3284 (AMU 2704), macho, proveniente del Cerro Socopé, cerca de 30 km
(por carretera) al SO de Guajiro, Municipio Mauroa, Estado Falcén, Venezuela, cerca de
1250 m, colectado por Abraham MuaAREs-URRUTIA, Leo YAGUA y Domingo DaaL, el 8 de
Febrero de 1996.

Paratopotipos. - EBRG 3285 (AMU 2706), hembra, ULABG 4112 (AMU 2702), macho, y
ULABG 4113-4114 (AMU 2701 y 2705), hembras, mismos datos que el holotipo.

Ejemplares referidos. - USNM 216804, [Estado] Falcôn, Cerro Socopé, 84 km. NW of
Carora, 1260 m; CIEZAH 385 (AMU 2700), hembra, mismos datos que el holotipo.

Diagnosis. — HEYER (1994) caracteriza, aunque con cierta reserva, el complejo de especies de
Leptodactylus podicipinus-wagneri por la ausencia de pliegues o en el mejor de los casos con un
par de pliegues dorsolaterales. Partiendo de esta definiciôn, Leptodactylus magistris se incluye
tentativamente como miembro de este complejo de especies. Esta nueva especie se distingue de
las especies incluidas por HEYER (1994) dentro del complejo podicipinus-wagneri por la
siguiente combinaciôn de caracteristicas: (1) hocico subovoide en vista dorsal; (2) longitud de
los dedos manuales 1 = 11 = IV < II; (3) cresta ulnar insinuada por una linea de espiculas
dispuestas laxamente; (4) piel dorsal con diminutas espiculas terminadas en puntas querati-
nizadas; (5) pliegues dorsolaterales ausentes; (6) disco ventral ausente; (7) espinas pectorales
ausentes; (8) puntas de los dedos manuales no dilatadas; (9) dos espinas grandes, puntiagudas,
queratinizadas en cada dedo pulgar manual; (10) brazos moderadamente hipertrofiados; (11)
dedos manuales con rebordes presentes en los dedos II y IIT; (12) linea labial clara con bordes
bien definidos y extendiéndose hasta por debajo del timpano; (13) pliegue tarsal débil, con
espiculas presentes a lo largo de la cresta del pliegue; (14) planta de las patas traseras con
diminutas espiculas queratinizadas.

Source : MNHN, Paris

MUARES-URRUTIA 115

Tab. 1. - Medidas (en mm) de los ejemplares holotipo y paratopotipos de Leptodactylus
magistris. LRC, longitud rostro-cloacal; LCB, longitud cabeza; ACB, ancho
cabeza; n, nümero de ejemplares.

Hembras (n = 4) Machos (n = 2)

EE Te eu

ue z Ambito de arr
CEA variacién ME
eständar eständar

Ambito de
variaciôn

38.6 + 7.55 27.9-45.1 39.05 + 0.07 39.0-39.1
13.6 + 1.13 12.8-14.9 11.55+0.21 11.4-11.7
15.1 + 1.27 14.4-16.6 13.15 + 0.07 13.1-13.2
19.2 + 1.13 18.4-20.5 17.9 + 0.14 17.8-18.0
20.4 & 1.08 19.5-21.6 18.35 + 0.63 17.9-18.8
23.2 + 0.98 22.6-24.4 20.55 + 1.62 19.4-21.7

2.9 + 0.32 2.7-3.3 2.8+ 0.14 2.7-2.9

De las especies del complejo de Leptodactylus podicipinus-wagneri registradas por HEYER
(1994) en Venezuela y norte de Colombia, L. magistris se diferencia de L. colombiensis,
leptodactyloides, nesiotus, pallidirostris, petersii, sabanensis y validus por tener todas ellas
pliegues dorsolaterales presentes (ausentes en L. magistris). La ünica especie sin pliegues
dorsolaterales, L. diedrus, se diferencia de L. magistris por tener la punta del hocico redon-
deado (subovoide en magistris) y la linea clara postero-labial indistinta o no discernible
(visible, con bordes superior e inferior bien definidos, en magistris).

Descripciôn del holotipo. — Cabeza ligeramente mäs ancha que larga; espacio interorbital liso,
ligeramente convexo; distancia interorbital 2.0 veces mayor que la anchura del pârpado
superior; canto rostral no muy bien definido; regiôn loreal ligeramente céncava, inclinada;
labios no ensanchados; narinas elevadas, dirigidas lateralmente y ligeramente hacia aträs;
narinas ms cerca de la punta del hocico que del borde anterior del ojo; hocico subovoide en
vista dorsal; punta del hocico redondeado en vista dorsal, subovoide en vista lateral; diâmetro
del ojo ligeramente inferior a la distancia ojo-narina; distancia internarinal casi igual a la
distancia ojo-narina; timpano bien visible, con bordes ligeramente elevados, parcialmente
queratinizados; timpano separado del ojo algo mäs de 1.8 veces su diämetro horizontal;
pliegue supratimpänico presente, apenas cubriendo una pequeña porciôn del margen superior

Source : MNHN, Paris

116 ALYTES 15 (3)

Fig. 1. (a) Vista dorsal de la hembra (izquierda, paratopotipo EBRG 3284) y del macho (derecha,
holotipo EBRG 3285), y (b) vista ventral con igual orden de Leptodactylus magistris.

del timpano; lengua mâs larga que ancha, 1/3 posterior no adherido al piso de la boca; coanas
redondeadas, no cubiertas por repliegue palatal del arco maxilar; dientes vomerianos presen-
tes, dispuestos moderadamente por deträs y entre las coanas, alineados uno al lado del otro;
maxilar y premaxilar dentados.

Dorso liso con numerosas espiculas queratinizadas, diminutas; extremo posterior del
dorso ligeramente tuberculado; pliegues dorsales y dorsolaterales ausentes; vientre finamente
rugoso; piel del pecho y de la garganta lisa; brazo y antebrazo igual que el dorso; una linea
conspicua de espiculas recorriendo a lo largo el borde ventrolateral del brazo y antebrazo;
tubérculo ténar ausente; tubéreulo palmar entre redondeado y ligeramente cordiforme, con
bordes inconspicuos; tubérculos supernumerarios ausentes; tubérculos subarticulares gran-
des, muy elevados, redondeados vistos desde arriba, ovalados en vista lateral; puntas de los

Source : MNHN, Paris

MUARES-URRUTIA 117

dedos no terminadas en almohadillas; ültima falange homogéneamente ancha en toda su
longitud; dedos manuales libres; rebordes carnosos presentes en los dedos IT y III, reborde
algo mäs pequeño presente del lado interno de los dedos I y IV; primer dedo de igual longitud
que el segundo; cada pulgar con un par de espinas queratinizadas, negras; espinas subiguales,
tamaño moderadamente grande; brazos moderadamente hipertrofiados.

Abertura cloacal a nivel de la linea media de los muslos, dirigida posteroventralmente,
cubierta por un pliegue corto, con borde crenulado; abertura cloacal rodeada por estrias
relativamente profundas y anchas, dando un aspecto radiado a la regiôn cloacal; piel de los
muslos, pantorrillas y tarsos moderadamente rugosa, con numerosas espiculas dorsalmente,
lisa ventralmente; pliegue tarsal presente, semejante a una quilla, con numerosas espiculas
sobre y a lo largo del pliegue; pliegue tarsal se extiende desde la base del tubérculo
metatärsico interno hasta casi 2/3 del tarso, haciendo un recorriendo ligeramente oblicuo;
tubérculo metatärsico externo casi cuadrado, elevado, subcénico en vista lateral; tubérculo
metatärsico interno muy alargado, casi 2.8 veces mäs largo que ancho, cerca del doble del
tamaño del tubérculo externo; tubérculos supernumerarios ausentes; espiculas presentes en la
palma de la pata, numerosas; tubérculos subarticulares grandes, entre redondeados y ovala-
dos vistos desde arriba, elevados, entre ovoides y subcénicos en vista lateral; dedos pedales
con palmeadura basal; férmula de la palmeadura varia entre la pata derecha y la izquierda: I
1.0—1.0 II 1.0—(0.5-1.0) III 0.5 — 1.0" IV (0.5-1.0) —(1.0-1.0*) V; dedos pedales con pliegues
carnosos bien desarrollados; un reborde visible a lo largo del margen externo del quinto dedo,
desde la punta hasta la base del primer tubérculo subarticular; extremo distal de los dedos
pedales de igual ancho que la falange adyacente; talones se superponen cuando los muslos son
colocados en linea recta respecto al eje longitudinal del cuerpo; talôn alcanza entre el timpano
y el ojo cuando las patas traseras son extendidas hacia adelante.

Coloracién en vida (datos tomados en el campo, incluyendo variacién individual). —
Castaño muy oscuro dorsalmente con algunas manchas castaño negrusco redondeadas
(aspecto “leopardino”). Dorso de muslos, tibias y tarsos de igual color con bandas negras
transversales, unas bien definidas y otras algo difusas, intercaladas unas con otras; miembro
anterior dorsalmente igual que posteriores. Vientre crema con algunas manchas pardas que se
hacen mäs densas anteriormente formando une “red” que puede extenderse sobre el pecho y
parte posterior de la garganta; regiôn gular entre castaño y castaño oscuro, con manchas
blancas; labio inferior castaño oscuro con manchas blancas; labio superior con bandas
castaño oscuro (dos lacrimales, una postero-ocular) y dorado castaño (en la punta del
hocico); banda interocular castaño dorado con bordes negros. Pupila negra, redondeada; iris
castaño pälido no reticulado.

Coloracién en preservativo del holotipo. - Dorso castaño oscuro sin manchas evidentes;
cara dorsal de los brazos, muslos, tibias y tarsos con bandas transversales negras; banda
interocular clara algo difusa; cabeza lateralmente con dos lineas blancas bien definidas (una
desde el borde posterior del ojo hasta la comisura de la boca y otra desde el borde inferior del
ojo hasta el borde del labio superior) y otras lineas claras, indistintas, en la regiôn loreal y la
punta del hocico. Labio inferior castaño oscuro con lunares pequeños, blancos; garganta
castaño oscuro sin manchas, ligeramente mâs pälido que en el labio inferior; pecho crema-
castaño inmaculado; mitad anterior del vientre crema con manchas y lineas irregulares
castaño pälido, formando una suerte de entramado laxo; mitad posterior del vientre y cara

Source : MNHN, Paris

118 ALYTES 15 (3)

ventral de los muslos crema inmaculado. Cara posterior de los muslos castaño oscuro con una
linea clara, oblicua desde algo lateralmente a la regiôn cloacal hasta un poco antes de alcanzar
la coyuntura tibio-femoral conectando con el color crema ventral.

Medidas (en mm) del holotipo. - Longitud rostro-cloacal 39.0; longitud cabeza 11.4; ancho
cabeza 13.2; fémur 17.8; tibia 17.9; timpano 2.9; longitud ojo-punta hocico 6.6; longitud
ojo-narina 3.6; diâmetro horizontal del ojo 3.1; distancia interorbital 4.1; distancia interna-
rinal 3.3; ancho pärpado superior 2.1; pata posterior 19.4. Las medidas de los restantes
ejemplares paratopotipos aparecen en la tab. 1.

Variaciones. — Los restantes cinco ejemplares paratopotipos, cuatro hembras y un macho,
todos adultos, mostraron una apariencia general muy homogénea entre si y con el ejemplar
holotipo. Todos los animales exhiben la linea pälida postero-labial bien visible y definidos sus
bordes, labio inferior pardo oscuro con manchas redondeadas blancas, regiôn gular castaño
mäs pälido, extendiéndose hasta el pecho y porcién anterior del vientre, observändose en esta
parte (y, ocasionalmente, también en el pecho) un marmoreado con manchas blancuzcas; no
se insinüan pliegues dorsolaterales. Se detectaron diferencias exclusivamente en las siguientes
caracteristicas: el primer dedo manual en las hembras siempre es visiblemente mäs largo que
el segundo (en los dos machos adultos son iguales); una hembra (EBRG 3285) tiene la punta
del hocico casi truncado (en vista dorsal) y el labio inferior no muestra el patrôn descrito
anteriormente, sino mäs bien crema pälido; las hembras no muestran el brazo hipertrofiado
como los machos.

Historia natural y distribuciôn. — Leptodactylus magistris fue colectado en el remanso de una
pequeña quebrada, al lado de un camino de tierra. Los ejemplares fueron colectados en un
ärea relativamente abierta y con abundantes plantas herbâceas y arbustos muy pequeños pero
densos, dentro de una sucesién secundaria de bosque nublado. En ese momento, aproxima-
damente las 16.30 h, se escucharon cantos de este especie desde dentro del agua y entre las
densas hierbas y arbustos en la orilla del remanso. También se observaron varios nidos de
espuma dispuestos en la orilla, generalmente disimulados bajo grupos moderadamente
espesos de hierbas, o asociados a rocas semi-sumergidas rodeadas o cubiertas con alguna
vegetaciôn. Se colecté un ejemplar pequeño, presuntamente juvenil (CIEZAH 386, longitud
rostro-cloacal 18.7 mm). No se observaron parejas en amplexo; sin embargo, se encontraron
dos renacuajos pardo oscuros (posteriormente perdidos), similares a los ya descritos para
especies de este grupo de especies (MAxsON & HEYER, 1988; HEYER, 1994). De los ejemplares
examinados, una sola hembra (EBRG 3285) tenia la cavidad abdominal Ilena de huevos
maduros en los oviductos, de tamaño moderado (1.1-1.4 mm de diâmetro), bicoloreados
(negro-blanco), redondeados, frecuentemente con algunos lados achatados; las restantes tres
hembras adultas mostraron los oviductos convolutos, Ilenos de huevos diminutos (inmadu-
ros), blancos. Ambos machos adultos presentan los cuerpos grasos bastante voluminosos
(casi Ilenan toda la cavidad abdominal), e igualmente un buen desarrollo del par de espinas del
primer dedo manual y las hendiduras vocales (vocal slits) bien evidentes y completamente
abiertas.

En la misma localidad se encontraron un adulto de Hyla crepitans, activa a esa hora del
dia, y adultos de Mannophryne sp., con machos cantando. Estos ültimos no se encontraban
junto con los Leptodactylus, sino algo mäs retirados del borde del camino, en una ärea de
vegetaciôn algo mäs densa.

Source : MNHN, Paris

MUARES-URRUTIA 119

Fig. 2. - Mapa mostrando la localidad aproximada (asterisco) de colecta de los ejemplares de la serie
tipica de Leptodactylus magistris, en Cerro Socopé, Estado Falcôn, Venezuela. El ârea punteada
representa la cota de 1200 m. Los nümeros señalados por triängulos indican los poblados de (1)
Guajiro, y (2) Rancho Lara. COL, Colombia; ANT. HOL., Antillas Holandesas.

Hasta el presente, Leptodactylus magistris se conoce exclusivamente de la localidad tipica
(fig. 2). Sin embargo, no se descarta su presencia en sectores mâs bajos de Cerro Socop6 y en
montañas vecinas como Cerro Azul (cerca 1880 m) y Cerro Cerrôn (cerca 2080 m) que, junto
con Cerro Socopé, constituyen la Serrania de Ziruma. Estas montañas estän conectadas por

Source : MNHN, Paris

120 ALYTES 15 (3)

valles relativamente altos (mäs de 500 m), con condiciones climäticas, tipo y densidad de
cobertura vegetal y disponibilidad de agua comparables (Robert WINGFIELD, comunicaciôn
personal).

Etimologia. — El epiteto especifico en latin magistris (nombre plural del modo dativo,
significando “a [mis] maestros””) hace honor a mis tres profesores: Pascual SORIANO, Enrique
LA MARCA y Alexis ARENDS, cuya generosa amistad, paciente guia, y oportunos consejos me
han inspirado y permitido avanzar para alcanzar metas profesionales y personales que
parecian imposibles. A ellos, mi muy sentido agradecimiento.

RESUMEN

Se describe una nueva especie de rana silbadora del género Leptodactylus, L. magistris,
proveniente del Cerro Socop6, cerca 30 km. (por carretera) al SO de Guajiro, Municipio
Mauroa, Estado Falcén, oeste de Venezuela. Esta especie, tentativamente considerada miem-
bro del complejo de especies de L. podicipinus-wagneri, se distingue de los miembros de esta
agrupacién por la ausencia de pliegues dorsolaterales, la punta del hocico subovoide en los
machos y redondeada-semitruncada en las hembras, y una linea pälida postero-labial visible
y bien definida.

AGRADECIMIENTOS

Deseo agradecer a Domingo DaaL y Leo YAGUA por su ayuda en el campo, y a Alexis ARENDS por
su apoyo logistico y moral. Este trabajo forma parte del proyecto de Inventario Herpetolägico del Estado
Falcôn (N° S1-9112-030), dirigido por Alexis ARENDS, con financiamiento parcial por Fundacite-Falcôn.
Agradezco también a Enrique La MARCA quien gentilmente permitié el estudio de animales y el uso de
equipos de la Colecciôn ULABG bajo su cuidado. E. La MARCA y W. R. HEvER hicieron algunos valiosos
comentarios sobre este manuscrit.

LITERATURA CITADA

Hever, W.R., 1994. — Variation within the Leptodactylus podicipinus-wagneri complex of frogs (Amphi-
bia: Leptodactylidae). Smiths. Contrib. Zool., 546: 1-124.

La Marca, E. 1992. - Catlogo taxonémico, biogeogräfico y bibliogräfico de las ranas de Venezuela.
Univ. Los Andes Cuad. Geogr, 9: 1-197.

== 1994. - Taxonomy of the frogs of the genus Manmophryne (Amphibia: Anura: Dendrobatidae). Publ.
Amigos de Doñana, 4: 1-15.

Maxson, L. R. & HEvER, W. R., 1988. — Molecular systematics of the frog genus Leptodactylus
(Amphibia: Leptodactylidae). Fieldiana-Zool., (n.s.), 41: 1-13.

RiveRo, J. À. 1961. - Salientia of Venezuela. Bull. Mus. comp. Zool., 126 (1): 1-207.

Corresponding editor: W. Ronald HEER.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 15 (3): 121-126. 121

The tadpole of Atelognathus nitoi
(Leptodactylidae, Telmatobiinae)

Néstor G. Basso * & Carmen A. UBEDA **

* Instituto de Limnologia “Dr. Raül A. Ringuelet”, Casilla de Correo 712, 1900 La Plata,
Argentina
** Departamento de Zoologfa, Centro Regional Bariloche,Universidad Nacional del Comahue,
Unidad Postal Universidad, 8400 Bariloche, Argentina

The tadpole of Atelognathus nitoi is described. Tadpoles of this
isolated species, endemic to the Nahuel Huapi National Park (Argentinian
Patagonia), show the basic characteristics of the described tadpoles in the

patagonicus and À. reverberii. Tadpoles of A. nitoi are more
similar in their external features and proportions to the tadpoles of À.
patagonicus but are easily distinguishable from both species by having a
broadly rounded tip of the tail and a smaller lateral emargination of the oral
disc. Further differences with A. reverberii are the smaller size, the more
slender body and the smaller oral disc.

INTRODUCTION

The genus Atelognathus consists of seven endemic species with small ranges in Patagonia
(southern South America). These species are mainly distributed around basaltic lagoons of
the extra Andean highlands in the provinces of Neuquén (4. patagonicus, À. praebasalticus),
Rio Negro (4. reverberii, À. solitarius) and Chubut (4. salai) in Argentina, between 38°40/ and
46°10'S, and in the Magellanic moorlands of Wellington Island (4. grandisonae) in southern
Chile, at about 49°S (Cet, 1984). Arelognathus nitoi inhabits a different habitat near a small
Andean lagoon surrounded by the deciduous temperate forest of Norhofagus pumilio in
western region of Rio Negro Province.

Only the tadpoles of A. patagonicus and A. reverberii have been described. These
nongregarious, free-swimming tadpoles of shallow waters have a typical leptodactylid appear-

ance. Tadpoles of 4. reverberii are larger and stouter than are those of 4. patagonicus (CEI,
1965, 1969).

While conducting field work in December 1995 and March 1996 at the type locality of 4.
nitoi (BARRIO, 1973), we collected tadpoles of this poorly known species. The description of
these tadpolesis here provided.

Source : MNHN, Paris

122 ALYTES 15 (3)

N
&
ARGENTINA e,
%
Xe
©
£
È
Refugio Neumeyer
Ê
Laguna Verde o 2
e
o
Cerro Chalhuaco &
2060 m
A Cerro Blanco
28m 0 1 2Km
nn —
7118. W

Fig. 1. - Location of Laguna Verde, type locality of Atelognathus nitoi, in the Parque Nacional Nahuel
Huapi, Rio Negro Province, Argentina. Triangle represents the peak of Cerro Challhuaco and
house symbol represents Neumeyer refuge.

MATERIAL AND METHODS

Tadpoles of Atelognathus nitoi were collected at the type locality: Laguna Verde, a small
lagoon on Cerro Challhuaco, Nahuel Huapi National Park, Rio Negro Province, Argentina
(41165, 71°18 W; ca. 1550 m of elevation), the only known locality of the species (fig. 1).
Geographic coordinates were obtained with a Garmin 45 Global Positioning System
(GPS).

Source : MNHN, Paris

Basso & UBEDA 123

10mm

Fig. 2. - Lateral view of a tadpole of Arelognathus nitoi (MACN 36695, stage 34).

Tadpoles deposited at Museo de La Plata (MLP A.1223) were collected on 10 December
1995 at developmental stages 25-28 (GOsNER, 1960). Some tadpoles were raised in the
laboratory and fixed at various stages of development. Three tadpoles were allowed to
complete metamorphosis in order to verify their specific identification. A tadpole deposited at
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN 36695) was col-
lected on 2 March 1996 at stage 34. AIl tadpoles were fixed in 10 % neutral formalin.
Measurements were taken with a dial caliper to the nearest 0.05 mm. Drawings were made
with a camera lucida attached to a Wild M-5 stereoscopic microscope. Terminology follows
ALTIG (1970), ALTIG & JOHNSTON (1989), Dugois (1995) and VAN Duxk (1966).

Specimens examined for comparisons included 4. patagonicus (MLP A.1222; 3 speci-
mens) and À. reverberii (MACN 28467-68; 2 specimens).

DESCRIPTION OF TADPOLES OF ATELOGNATHUS NITOI
(fig. 2-3)

Type IV tadpoles (ORTON, 1953). Total length of largest specimen of the series (stage 40),
56.45 mm; smallest tadpole (stage 25), 15.1 mm (tab. 1). Total length at stage 34 (MACN
36695), 47.35 mm; body length about two-fifths of total length, almost twice body width;
body shape oval, somewhat depressed, 1.2 times wider than high; snout rounded in dorsal and
lateral profile; eyes large, dorsolateral; interorbital distance twice eye diameter; nostrils
dorsal, protuberant, closer to eye than snout; distance between nostrils equal to their distance
from eyes and three-fifths of interorbital distance; spiracle sinistral, opening directed poste-
rolaterally; vent tube short; vent opening dextral, lying dorsal to a fold in ventral fin; caudal
musculature moderately robust anteriorly, becoming narrower gradually toward tip of tail;
dorsal and ventral fins equally developed, higher than musculature at midlength; dorsal fin
extending slightly onto body; tip of tail rounded.

Oral disc anteroventral, slightly wider than interorbital distance, about 40 % of body
width; one row of small regular marginal papillae, with wide medial gap on upper labium (fig.
3); disc emarginate, lateral margin with small, but distinct constrictions; intramarginal lateral
papillae present in infra- and supraangular regions; tooth row formula 2[A;]/3[P,]; upper
rows as long as upper sheath; third lower row slightly shorter than second lower row; jaw
sheaths thin, gently curved and finely serrate.

Source : MNHN, Paris

124 ALYTES 15 (3)

Tab. 1. - Measurements (mm, mean + standard deviation) of tadpoles of Atelognathus nitoi.
Those marked with an asterix (*) were raised in the laboratory from tadpoles
collected at stages 25-28. n: sample size.

Body length Total length

7.28 + 0.54 15.76 + 1.20
8.22 + 0.48 17.74 + 0.91
9.64 + 0.40 20.54 + 0.86
10.54 + 0.50 22.76 + 2.68
14.05 39.50
21875 47.35
17.62 + 1.08 42.99 + 1.29
21.42 + 0.25 51.32+2.37
22.25 54.65
22.63 + 0.47 54.62 + 2.33

C7
OUT 5
OUTTrE Ce
CPL DPI

D : g
CUT
MAT

CET
772

C7 v
CET PP PTE us

mm

Fig. 3. — Oral disc of a tadpole of Atelognathus nitoi (MACN 36695, stage 34).

Source : MNHN, Paris

Basso & UBEDA 125

In life, dorsal color pattern of body golden-brown with bilateral unpigmented areas
posterior to gills, giving the appearance of an indentation between head and body; caudal
musculature golden-brown; caudal fins transparent with dark markings. In ventral view,
intestine, heart, and cranial structures visible. In preserved specimens, densely crowded
melanophores on the posterior body mark courses of blood vessels in tail musculature and
fins.

DISCUSSION AND CONCLUSIONS

Tadpoles raised in the laboratory were maintained on a diet of Tetra Phill fish food. The
tadpoles collected on 10 December 1995 and raised through metamorphosis in the laboratory
reached stage 46 between 18 February and 11 March 1996.

The tadpoles collected at stages 25-28 were found at the littoral zone of the lagoon where
they spent a considerable amount of time on the bottom in the shallow water. When
disturbed, they swam individually about 1 m to a different location. The tadpole at stage 34
was found buried in the fine sediment on the bottom of the lagoon in 3 m of water. One adult
female was found under a rotten tree-trunk near the pond margin on 10 December 1995.

The tadpoles of 4. nitoishow the basic diagnostic characters listed by LAVILLA (1988) for
the genus Atelognathus and can be classified in the ecomorphological guild Benthic (Section
11.12) of ALTIG & JOHNSTON (1989).

Of the known tadpoles of the frog genus Atelognathus, tadpole of 4. nitoi is most similar
to that of À. patagonicus (Cet, 1965) from Laguna Blanca (Neuquén Province) than to that of
A. reverberii (Cr1, 1969) from Somuncurä Plateau (Rio Negro Province). The tadpole of 4.
nitoi may be easily distinguished from À. patagonicus and À. reverberii by the broadly rounded
tip of the tail and the smaller emargination of the oral disc. It differs further from the tadpole
of À. reverberii by being smaller (4. reverberii grows to 98 mm total length; Cet, 1969), by
having a slender oval body (4. reverberii has a stout and depressed body), and by having a
smaller oral disc.

RESUMEN

Se describe la larva de Atelognathus nitoi en base a especimenes colectados en la localidad
tipo de la especie: Laguna Verde, Cerro Challhuaco, Parque Nacional Nahuel Huapi,
Provincia de Rio Negro, Argentina. Las larvas de 4. nitoi presentan las caracteristicas tipicas
descriptas para el género. De las dos especies cuyos renacuajos se conocen, À. nitoi presenta
larvas con una morfologia mas similar a la larva de À. patagonicus que a la de À. reverberii.
Las principales caracteristicas externas diferenciales con À. patagonicus y À. reverberii son el
extremo caudal notablemente redondeado y las emarginaciones laterales del disco oral menos
pronunciadas. De 4. reverberii se diferencia, ademäs, por el menor tamaño general, el cuerpo
menos deprimido y el disco oral relativamente mas chico.

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126 ALYTES 15 (3)

ACKNOWLEDGMENTS

We thank Gustavo CarRizO, from the Museo Argentino de Ciencias Naturales “Bernardino
Rivadavia”, for provision of working space and loan of specimens for comparisons. The Administraciôn
de Parques Nacionales of Argentina allowed us to develop field studies at the Nahuel Huapi National
Park. Financial support for fieldwork was provided in part by a grant from the Eppley Foundation for
Research to Juan À. SCHNACK and Gustavo R. SPINELLI for the project entitled “Biodiversity in
Patagonia”. We are grateful to Robert F. INGER, Maureen A. DONNELLY and an anonymous reviewer for
helpful suggestions on the manuscript. This study was partially supported by an external fellowship from
the Consejo Nacional de Investigaciones Cientificas y Técnicas of Argentina to NGB and a grant from
the Universidad Nacional del Comahue to CAU.

LITERATURE CITED

ALTIG, R., 1970. — A key to the tadpoles of the continental United States and Canada. Herpetologica, 26:
180-207.

ALriG, R. & Jonnsron, G. F., 1989. - Guilds of anuran larvae: relationships among developmental
modes, morphologies, and habitats. Herp. Mon., 3: 81-109.

BaRRI0, À. 1973. - Una nueva especie de Telmatobius (Anura, Leptodactylidae) procedente del dominio
austral cordillerano argentino. Physis, 32 (84): 207-213.

Cri, J. M. 1965. - The tadpole of Batrachophrynus patagonicus Gallardo. Herpetologica, 20: 242-245.

ee 1969. The Patagonian telmatobiid fauna of the volcanic Somuncura plateau of Argentina. J Herp.,

3: 1-18.
1984. — A new leptodactylid frog, genus Atelognathus, from southern Patagonia, Argentina. Herpe-
tologica, 40: 47-51.

Dusois, A., 1995. - Keratodont formulae in anuran tadpoles: proposal for a standardization. J zoo. Syst.
Evol. Res., 33: I-XV.

Gosner, K. L., 1960. — A simplified table for staging anuran embryos and larvae with notes on
identification. Herpetologica, 16: 183-190.

LaviLLA, E. O., 1988. - Lower Telmatobiinae (Anura: Leptodactylidae): generic diagnoses based on
larval characters. Occ. Pap. Mus. nat. Hist. Univ. Kansas, 124: 1-19.

ORTON, G. L., 1953. - The systematics of vertebrate larvae. Syst. Zool., 2: 63-75.

VaN Dux, E. D., 1966. - Systematic and field keys to the families, genera and described species of
southern African anuran tadpoles. Ann. Natal Mus., 18: 231-286.

Corresponding editor: Ronald G. ALTIG.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 15 (3): 127-132. 127

A redescription
of Ramanella mormorata Rao, 1937
(Anura, Microhylidae)

Indraneil Das & Romulus WHITAKER

Centre for Herpetology, Madras Crocodile Bank Trust, Post Bag 4,
Mamallapuram, Tamil Nadu 603 104, India

Ramanella mormorata Rao, 1937, hitherto known only from the
suntypes collected over half a century ago from “Saklespur, Hassan District,
Mysore”, at present in Karnataka State (south-western India), that are now
lost, is redescribed, based on a new specimen from Cotigao Wildli
Sanctuary (Goa, south-western India), and an older one from “Malabar”
the collection of the Museum of Comparative Zoology. The species is
compared with congeneric species from southern India and Sri Lanka.

INTRODUCTION

The genus Ramanella was described by RAO and RAMANNA (1925), and named for the
junior author, the type species (by monotypy) being Ramanella symbioitca Rao & Ramanna,
1925 (apparently a misprint for Ramanella symbiotica), a name considered a junior syn-
onym of Callula variegata Stoliczka, 1872 by PARKER (1934: 93) and FRosT (1985: 389). The
genus, redefined by PARKER (1934), is restricted to peninsular India and Sri Lanka, and
eight nominal species are currently recognized, five from India, two from Sri Lanka and one
from both regions (see FRosT, 1985: 389). Of these, perhaps the least known are three spe-
cies from the Western Ghats of south-western India, Ramanella anamalaiensis, Ramanella
minor and Ramanella mormorata, that are known only from the types described sixty years
ago by RAO (1937) and then deposited in the Central College Museum in Bangalore. Since
then, the whole collection of specimens described by RAO (1937) has been lost (see DUBoIS,
1984: 156-157). The present paper is devoted to one of these species, R. mormorata, the lost
type specimens of which were stated to be from “Saklespur, Hassan District, Mysore”
(currently spelt “Sakleshpur”; 12°59/N 75°43’E; at present in Karnataka State, south-western
India).

When he prepared the original description of Ramanella mormorata, although the
number of types was not specified, RAO (1937) clearly had several specimens of his new
species, including “young specimens”, “immature specimens”, “mature males” and “mature
females”, but he presented measurements of a single specimen. As no holotype was desig-

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128 ALYTES 15 (3)

nated in this original description, all these specimens must be considered syntypes of this
species. RAO (1937: 420) stated that these specimens came from “whorls of the plantain
leaves”, and were usually found solitary, although occasionally two or three could be found
together.

The purpose of this note is to report the rediscovery of Ramanella mormorata, based on
material from Goa (India), a distance of over 300 km north-west of the type locality, and the
discovery of an older specimen from “Malabar” in the collection of the Museum of Compar-
ative Zoology (MCZ, Cambridge, USA). The species is redescribed and compared with the
original description in RAO (1937).

One specimen (MCZ A.116283) was collected by the second author on 26-28 October
1994 from a forest rest house in Canacona (15°01°N 74°04’E), adjacent to Cotigao Wildlife
Sanctuary (Goa, south-western India). This collection locality lies contiguous to a patch of
wet evergreen forest, in a lowland situation. A second specimen (MCZ A.15421) was found in
the MCZ collection with the locality “Malabar”, which refers generally to the entire south-
western highlands of India, also known as the Western Ghats.

MATERIAL AND METHODS

Measurements (to the nearest 0.1 mm) were taken with a Mitutoyo dial vernier calliper
from specimens preserved in 70 %ethanol, over 12 months after collection in the case of MCZ
A.116283. The following measurements were taken: SVL, snout-vent length (from tip of
snout to vent); TBL, tibia length (distance between surface of knee and surface of heel, with
both tibia and tarsus flexed); A-G, axilla to groin (distance between posterior edge of fore
limb at its insertion to body and anterior edge of hindimb at its insertion to body); HL, head
length (distance between angle of jaws and snout-tip); HW, head width (measured at angle of
jaws); HD, head depth (greatest transverse depth of head, taken at the orbital region); ED, eye
diameter (diameter of orbit); UE, upper eyelid width (greatest width of upper eyelid); IO,
interorbital distance (least distance between upper eyelids); IN, internarial distance (distance
between nostrils); E-S, eye to snout-tip distance (distance between anteriormost point of eye
and snout-tip); E-N, eye to nostril distance (distance between anteriormost point of eye and
nostril); and F2D, diameter of the disk on finger II.

DESCRIPTION

A small Ramanella (SVL up to 29.6 mm); habitus robust; body ovoid, with a thick waist
(fig. 1-2); head short (HL/SVL ratio 0.20-0.24), broad (HW/SVL ratio 0.30-0.33); snout
obtuse when viewed dorsally, truncate in lateral view, in level with the mandible, nostrils closer
to tip of snout than to eyes (E-N/E-S ratio 0.54-0.56); canthus rostralis sloping; loreal region
oblique; eyes large (ED/HL ratio 0.47-0.49), eye diameter greater than eye-nostril diameter
(ED/E-N ratio 1.45-1.53); interorbital distance over two-and-half times greater than upper
eyelid width (IO/UE ratio 2.27-3.07); distinct occipital fold (fig. 3); pupil vertical; supratym-
panic fold distinct, extending from posterior corner of upper eyelid to insertion of forelimb at

Source : MNHN, Paris

Das & WHITAKER 129

1.1 12:

Fig. 1. - Ramanella mormorata from Canacona, near Cotigao Wildlife Sanctuary, Goa, south-western
India (MCZ A.116283) in dorsal (1.1) and ventral (1.2) views. Bars: 10 mm.

2.1 2,2

Fig. 2. - Ramanella mormorata from “Malabar”, south-western India (MCZ A.15421) in dorsal (2.1) and
ventral (2.2) views. Bars: 10 mm.

Source : MNHN, Paris

130 ALYTES 15 (3)

Tab. 1. - Measurements (in mm) of Ramanella mormorata Rao, 1937 from south-
western India (see text for details).

Measurements MCZ A.15421 MCZ A.116283

Snout-vent length 29.6 25.9
Axilla-groin distance 13.4 11.0
Head length 5.9 6.2
Head width 9.0 8.6
Head depth 5-5) 5.0
Eye diameter 29 2:9
Upper eyelid width 15 1.4
Interorbital distance 3.4 3.4
Internarial distance 1.8 2.4
Eye-snout-tip distance 34 3.4
Eye-nostril distance 2.0 1.9
Tibia length 1127 10.8

Diameter of disk on finger II 172; 12

Fig. 3.- Forehead of Ramanella mormorata (MCZ A.15421), showing the distinct occipital fold.

Source : MNHN, Paris

Das & WHITAKER 131

axilla; tympanum not externally visible; nostrils laterally oriented; inferior aspect of snout-tip
smooth, without any nicks; inner margin of mandible with a distinct W-shaped notch; tongue
large, smooth, without papillae, oval, measuring 6.2 mm in greatest length (in MCZ
A.116283); post-choanal ridge well developed, with a broad median gap; cloacal opening
directed posteriorly, slightly below the upper level of thighs.

Forelimbs long, the tips of fingers dilated into triangular, flattened disks, the largest on
finger II (measuring 1.2 mm in both specimens examined), lacking webbing; two elongated
metacarpal tubercles. Relative length of fingers: 3 > 4 > 2 > 1. Hindimbs relatively short; tibia
short (TBL/SVL ratio 0.40-0.42); tibio-tarsal articulation failing to reach axilla; heels failing
to overlap when the hindimbs are set at right angles to the body; tips of toes weakly swollen,
but not dilated; toe webbing vestigeal, with three phalanges free on toe IV; a large elongated
inner metatarsal tubercle and a smaller rounded outer metatarsal tubercle. Relative length of
toes:4>3>5>2> 1. Measurements are given in tab. 1.

Skin with large, flattened pustules scattered throughout the dorsum; ventrum smooth.

In preservative, dorsum of MCZ A.116283 pale olive with dark brown bands behind the
internarial, the interorbital and over the scapular region. Scattered dark brown blotches
present on the rest of the dorsum; fore- and hindimbs dark-barred, the most distinctive bars
being on the inguinal region, which is joined through the thighs to across the cloaca. Venter
pale yellow, variegated with dark brown on the throat, chest, abdomen and the undersurfaces
of the fore- and hindimbs. MCZ A.15421 is relatively more discoloured, but shows the
dark blotches on the dorsum and the limb bands; the dark bands on the dorsum are broken
up.

COMPARISONS

The present material matches the original description and illustrations of Ramanella
mormorata by RAO (1937: 419-420, pl. 29 fig. 19-19a) in the following characters: well
developed post-choanal ridges, with a broad median gap; toes webbing weak; dorsum with
dark blotches, in addition to dark bands behind the internarial, interorbital and scapular
regions; dark barred thighs that are fused, running along the thighs over the cloaca and up the
inguinal region. The specimen (sex unspecified) measured by RAO (1937: 419-420) was 25.0
mm in snout-vent length, while the two specimens being reported here (both females) measure
25.9 mm (MCZ A.116283) and 29.6 mm (MCZ A.15421).

However, the two new specimens do not fit the original description of RAO (1937) in a
couple of apparently fundamental characteristics. Both specimens show a distinct occipital
fold (see fig. 3), which was reported absent in RAO’s specimens, and they show a conspicuous
supratympanic fold, described as “inconspicuous”’ by RAO (1937). Since the new locality (in
Goa)is over 300 km from the type locality, such variation is not remarkable. In addition, many
of Rao’s (1937) types were clearly desiccated specimens, which may account for the damage
to dermal features. In Ramanella montana (Jerdon, 1854), Ramanella obscura (Günther,
1864), Ramanella variegata (Stoliczka, 1872) and Ramanella triangularis (Günther, 1876), the
occipital fold is occasionally absent (see PARKER, 1934).

Source : MNHN, Paris

132 ALYTES 15 (3)

Ramanella mormorata differs from all its congeners in coloration. Its small, dark blotches
and transverse bands contrast with the dark, median, hour-glass figure of R montana, R
obscura, R. palmata and R. triangularis, with the broad median, dark band of R. anamalaien-
sis, and with the broad, dark triangle on the anterior part of the dorsum of R. minor. R.
mormorata also differs from R. obscura, R. palmata, R. triangularis and R. variegata in having
the eye-snout distance greater than the diameter of the eye (vs. eye-snout equal to or less than
eye diameter in its congeners), well-developed post-choanal ridges (indistinct or absent in R.
anamalaiensis and R. variegata) and toes weakly webbed (vs. free in R. anamalaiensis and R.
minor, webbing on toe IV reaching the basal subarticular tubercle as a broad sheath and
continuing as a narrow sheath to the base of the swollen tips in R. montana, and up the distal
subarticular tubercle in R. palmata). R. mormorata differs from R. minor in showing interna-
rial distance greater than the upper eyelid width (vs. equal). It also differs from R. montana in
showing a broad median gap between the post-choanal ridges (vs. ridges nearly in contact)
and free fingers (vs. fingers distinctly webbed, fide PARKER, 1934) and from R. obscura in
having a supratympanic fold thin (vs. with a parotoid-like thickening).

ACKNOWLEDGEMENTS

Manuscrit preparation was supported by the Centre for Herpetology, Madras Crocodile Bank
Trust and a Fulbright Fellowship to the first author. We thank José Rosapo (MCZ), for curatorial
assistance, and Sushil DUTTA, Robert INGER and an anonymous reviewer for commenting on an earlier
draft of the manuscript.

LITERATURE CITED

Duroïs, À. 1984. - Note préliminaire sur le groupe de Rana limnocharis Gravenhorst, 1829 (Amphibiens,
Anoures). Alytes, 3 (4): 143-159.

FRosr, D. R. (ed.), 1985. - Amphibian species of the world. Lawrence, Allen Press & Ass. Syst. Coll.:[i-iv]
+iv + 1-732.

PARKER, H. W. 1934. — À monograph of the frogs of the family Microhylidae. London, British Museum
(Natural History): i-viii + 1-208.

R40, C. R. N,, 1937. — On some new forms of Batrachia from S, India. Proc. indian Acad. Sci., (B), 6:
387-427, pl. 21-31.

Ra, C. R. N, & RAMANNA, B.S., 1925. — On a new genus of the family Engystomatidae (Batrachia). Proc.
zool. Soc. London, 1925: 587-597.

Corresponding editors: Masafumi MATsuI & Alain DuBois.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 15 (3): 133-136. Book review

An evolutionary biologist’s view
on the science of biology

Alain DuBoIs

Laboratoire des Reptiles et Amphibiens,
Muséum national d'Histoire naturelle,
25 rue Cuvier, 75005 Paris, France.0

Mayr, Ernst. — This is biology. The science of the living world. Cambridge, Massachusetts & London,
England, The Belknap Press of Harvard University Press, 1997: i-xvii + 1-327.

Born in 1904, Ernst MAyR is now a living legend of evolutionary biology. His countless original
contributions to the development of this discipline were published in about 650 journal articles and about
twenty major books, starting with Systematics and the origin of species (1942), and going through
landmarks such as Animal species and evolution (1963), Principles of systematic zoology (1969) or The
growth of biological thought (1982). Ernst MaAYR's last book, This is biology (1997), his last brilliant
contribution to the understanding of the unique characteristics of biology among the sciences and of its
particular philosophical bases, is of particular importance for all biologists. It provides a detailed analysis
of several major questions often put by biologists, by scientists of other disciplines, and by lay persons,
about the science of the living world.

Much has been written about the history and epistemology of biology, but many of the authors of
these works were historians of science or philosophers, not biologists. The interest and significance of
Mayr’s new book come mostly from its having been written by a biologist, who has an inside under-
standing of the problems of this scientific field. As is very aptly shown by Mayr in this book, the
philosophy and history of science have long been dominated by a “conception of science” derived from
physics, and such a conception cannot address many aspects of the science of life. The characteristics of
biology are very different from those of physical sciences and other sciences of matter, and generalisations
drawn from the study of the latter often do not apply to biology, because of the unique particularities of
life among natural phenomena.

Among the important differences, physics and other matter sciences try and draw laws having a
general, universal value. À long-prevailing conception has also tried to assign this aim to biology. Such a
conception can be illustrated by sentences such as: “science only deals with general matters” or “if you
explain the bacterium to me, l'Il leave you quits with the elephant or man”. In such a perspective, it would
be enough to study in every detail a single kind of organisms (a bacterium, a fruit fly, a white mouse) to
know everything about life. However, this reductionist perspective has severe limits.

For sure, a number of particularities are common to all or most organisms, such as the genetic code,
the basic cellular biochemical and physiological processes, or certain characters common to members of
major clades. But, beside these common features, organisms are more correctly characterized by their
diversity and the diversity of their characters. Contrary to what the reductionist approach suggests, this
diversity is not a secondary phenomenon, or a disturbing “noise” in the study of life, but it is the main
characteristic of life. This diversity is the result of the evolution of organisms on our planet, which has
involved two distinct mechanisms: a progressive modification of the characteristics of organisms within
evolutionary lines (process of anagenesis), and a multiplication of the clades of organisms (process of
cladogenesis): both mechanisms together are traditionally known as the process of phylogenesis. While the
terms phylogenesis, anagenesis and cladogenesis refer to processes, their results may be known together as
phylogeny, and 1 further suggest that the components of the latter could be recognized separately as
anageny and cladogeny. As a consequence, biology is characterized much less by the existence of laws than

Source : MNHN, Paris

134 ALYTES 15 (3)

by the fact that most of the characteristics of organisms can be explained because these organisms are the
result of a history. Although this is not fully understood by many biologists, much more than a
deterministic science, biology is a historical science.

Another important particularity of biology, well underlined in MaYR's book, is that nothing makes
sense in biology if one does not understand that biological phenomena have several distinct levels of
integration. This means that the properties of an organism cannot be reduced to those of its cells or
organs, those of a population to those of its individuals, etc. At each level of integration, new particular-
ities emerge, wich cannot be mechanically deduced from the particularities present at the immediately
lower integrative level. This very old notion can be summarized in a sentence like: “the whole is more than
the sum of its parts”. It is qualified by Ernst MAYR, after others, under the term of emergence.
Consideration of this characteristic of life is of paramount importance to understand many biological
phenomena, and it supports a holistic, rather than a mechanistic or reductionist, approach to these
phenomena.

Another strong quality of Mayr’s book is to show in detail how, in science, several types of questions
can be legitimately asked, and how these questions can be reduced to three major categories: the questions
“what”, “how” and “why”. Many scientists, and even biologists, tend to believe that the only “legitimate”
scientific questions are “how” questions: how does such biochemical or physiological mechanism work,
how does things work inside the “black box” of such behaviour, etc. Such questions can often be studied
through an experimental approach, and, to this day, some biologists still tend to think that scientific
knowledge can be obtained only through experimental method. This is easy to understand in countries,
such as France, where biology has long been a mere subdiscipline of medicine, and where great historical
figures in research are those of people, such as Claude BERNARD or Louis PASTEUR, who were wearing
white gowns and devoted most of their life to laboratory experimental studies. The situation is different
in Anglo-Saxon countries. where there exists a strong tradition of field naturalists, such as DARWIN or
May, which has facilitated the emergence of a different, more holistic, approach to biology.

As a matter of fact, it is totally incorrect to state that the question “how” is the only legitimate
question in science. This question allows to elucidate only one type of problems in biology, those which
can be designated under the term “proximal causes”. Thus the experimental method allows to answer a
question such as “what is the environmental factor that determines the fact that an animal starts a
seasonal migration?”, but not the question “why did seasonal migrations appear in this species?”. Such
“why” questions aim at elucidating “evolutionary causes” of biological phenomena. They are as
legitimate as “how” questions, because in fact each biological phenomenon is the result of wo distinct
causation systems: evolutionary and proximal causes. More and more biologists have become aware of
this double causation system and recognize the legitimacy of “why” questions in science. Usually such
questions cannot be answered to through the experimental method. They are the major questions which
“evolutionary biology” asks. For a long time, the only answers to such questions were theological ones,
but since 1859 and the development of the concept of natural selection it has been possible to provide
scientific answers for them.

But there is a third kind of questions in science: these are “what” questions. Questions of this kind
are the first ones man asks in front of the world: what exists, what are the major characteristics of what
exists, etc. The answer to such questions requires to have recourse to observation, description, compari-
son or inventory. Contrary to an ideology currently dominant in science, such questions are legitimate
scientific questions. As long as no correct answer has been provided to them, the questions “how” and

why” are meaningless, or at least cannot be correctly set. What” questions must therefore be respected

and this also applies to those who study them. As a matter of fact, the dominant activity of scientists in
allscientific fields (including the most “modern” ones) are of this kind. Careful and complete descriptions
and inventories are the first steps that cannot be done without in all fields, including molecular biology
(description and inventory of molecules and of their activities), genetics or ecology.

An important proportion of “what” questions in biology is the set of questions that relate to the
inventory and classification of biodiversity. Given the importance of these questions in the previous
works of Ernst MAYR, it is not surprising that he devoted a nice chapter to this topic in his new book.
Once again, MAYR comes back in this book to the many problems related to the building of biological
classifications, which he had already discussed in several other important works (MAYR, 1969, 1974, 1981;
MayYr & AsHLOCK, 1993). He very persuasively shows how the different “schools” of macrotaxonomy,

Source : MNHN, Paris

Dugois 135

and in particular the two dominant ones in the recent years (cladistics and evolutionary or “Darwinian”
classification) do in fact differ in their objectives. The aim of an evolutionary classification is to provide a
taxonomy, i.e. a hierarchical arrangement of taxa. The latter are classes that should be recognized on the
bases of two criteria: genealogy (common descent) and degree of similarity (amount of evolutionary
change). These two criteria correspond to the two dimensions of phylogeny: anageny and cladogeny.
MayYr quite rightly suggests that arrangements of organisms built on the basis of cladogeny alone do not
deserve the qualification of true classifications, but should rather be known as cladifications. The latter
recognize units which are not classes (as are taxa), but clades, renamed by MAyR (1995) as cladons. To
complete MAyR’s terminological clarification, some other terms can be useful: the term phylon, first
proposed by Dusois (1991) to designate the concept later called cladon by Mayr (1995), would be more
appropriate to designate a phylogenetic s.L. i.e. both anagenetic and cladogenetic, unit. Following MAYR’s
conception of classification, the term phylon is therefore a strict synonym of the term taxon. While
classification according to phylogeny s., i.e. both anageny and cladogeny, gives birth to what can be
called either a faxonomy or a phylonomy, cladification, ie. classification based on cladogeny alone, results
in a cladonomy, not properly a taxonomy. The differences between the two systems can also be stressed
when one considers the way characters are used to define taxa. In a traditional taxonomy, a taxon can
validly be diagnosed, i.e. characterized by a diagnosis (or taxognosis, or phylognosis): this is a set of
differential or diagnostic characters, both plesiomorphic and apomorphic ones, that characterize this
taxon and distinguish it from related ones. On the other hand, in a cladonomy a cladon needs only be
apognosed, i.e. characterized by an apognosis (or cladognosis, or more shortly clagnosis): the latter only
includes apognostic characters, i.e. autapomorphic characters of the cladon, not shared with closely
related ones.

Such terminological discussions may appear gratuitous or superfluous to some, but they are not. As
shown on several occasions in MAYR's book, during the whole history of biology, many scientific debates,
discussions and conflicts turned out to be ultimately caused by terminological confusions. Many so-called
disagreements between colleagues simply take their root in the fact that these different biologists used the
same term in different senses. Introduction of the term cladification is therefore a particularly useful
contribution, which hopefully will be followed by all evolutionary biologists and systematists. It will help
more and more people to understand that both classification and cladification may be legitimate, but that
they do not have the same objectives. Evolutionary (or Darwinian, or synthetic) classifications serve
multiple purposes, both practical and theoretical: their aim is to provide a hierarchical arrangement of
taxa, the latter being non-polyphyletic and homogeneous groups of populations or taxa, about which the
highest possible number of generalisations and predictions can be made. On the other hand, the aim of
a Hennigian cladification is merely to give a transcription, under the form of a hierarchical arrangement,
of a cladogram, and therefore to provide information on the branching pattern of clades in the phylogeny
of a group. Both aims may be justified, depending on the information one wants to obtain, but it is
important not to confuse both kinds of information storing systems.

Another clarification is wanting in MAYR’s new book, as in his previous texts: MAYR rightly stressed
on several occasions that the term monophyletic was used by cladists for a concept quite different from
that designated by HAECKEL (1868) when he coined this term. As a result, in this book MAYR once again
claims that the term monophyletic (and its derivative monophyly) should be restored in their original
senses. Given the number of recent publications where these terms were used in HENNIG’s (1950) new
sense, I do not think this restoration will ever take place for all biologists, and, in my opinion, in order to
avoid the continuation of confusion, these terms should be abandoned altogether. For “monophyletic
sensu HENNIG” (i.e., a qualification of a group which is both non-polyphyletic and non-paraphyletic),
AsHLock’s (1971) term holophyletic should be used. For “monophyletic sensu HAECKEL” (non-
polyphyletic), Duois’s (1986) term homophyletic is available. My feeling is that in this case one should
follow the same line of reasoning as that advocated by MAyR & AsHLocKk (1991: 276-277) regarding the
case of the terms character, character state, signifer and signifer state: “We realize that the character-
character state terminology has been too widely adopted to be easily dislodged. Therefore, any endeavor
to restore the traditional meaning of the word character would cause considerable confusion. Hence,
although with considerable reluctance, we use character state for what traditionally has been called a
character.” A similar “considerable confusion” might arise from a continuous effort to restore the
traditional meaning of the term monophyletic, and much clarification would come from a rejection of
this term and its replacement by either holophyletic or homophyletic according to the purpose.

Source : MNHN, Paris

136 ALYTES 15 (3)

Mayr’s book is still rich of many other stimulating discussions and analyses. After “what”
questions, case studies of “how” and “why” questions allow to illustrate other important epistemological
aspects of biology as a science with no equivalent among other scientific fields. Two chapters are devoted
to the kind of questions that ecology asks, and to the status of our knowledge about human evolution.
Possibly the least convincing chapter is the last one, which deals with the relationship between our
knowledge of biological facts (and mostly evolution) and ethics. This chapter could have benefited from
consideration of other works published on this matter by authors not mentioned by MayR, such as the
numerous books of Jean ROSTAND, to give only one example. Strangely, regarding human phenomena,
May seems to share the conventional reductionist attitude of many other biologists, who think that
most psychological and social human features can be explained in terms of biology - if not simply of
“common sense” (see e.g. pp. 39-40, 254, 260, 264-265, 267). However, here also, emergence is at work. It
is as misleading to analyse these high-integration-level phenomena with biological concepts as to analyse
biological phenomena with concepts from biochemistry or physics. During the last century, all scientific
disciplines dealing with man (psychology, social sciences, economics, etc.) showed a great development
and experienced several “scientific revolutions” as significant as those of GALILEO and DARWIN in their
respective fields. Unless one is ready to accept that man, being “special among God's creations”, cannot
be studied scientifically, these developments must be duly considered. In order to adopt a scientific
attitude in this respect, there is the same need of terminological and conceptual clarity and rigour as that
aptly advocated by Mayr himself regarding biology, and, because scientists are humans, “common
sense” is even more misleading here as it is in biology. Ignorance by May of most of the significant
works and theories concerning human behaviour, psychology and society severely limits the interest and
reach of this chapter of his book.

No one now knows what the biology of the next century will be. Concerning the part of this science
that deals with the diversity of life (systematics), which has attracted most of the attention of MAYR in his
works, some authors (e.g.: CRowsoN, 1970; Duois, 1988) have suggested that “experimental systema-
tics”, dealing in particular with developmental problems, might be the next important step in this old
research field. Whatever the case may be, terminological and conceptual clarifications will be of
paramount importance for future progress of the discipline, and MaYR’s works will have provided a lot of
new and useful elements in this respect.

LITERATURE CITED

AsuLock, P. D., 1971. - Monophyly and associated terms. Syst. Zool., 20: 63-69.

CRowsoN, R. A., 1970. - Classification and biology. London, Heinemann: i-ix + 1-350.

Dusois, A., 1986. - À propos de l'emploi controversé du terme “monophylétique”: nouvelles proposi-
tions. Bull. Soc. linn. Lyon, 55: 248-254.

so 1988. — The genus in zoology: a contribution to the theory of evolutionary systematics. Mém. Mus.
nat, Hist, nat., (A), 140: 1-123.

—— 1991. - Nomenclature of parthenogenetic, gynogenetic and “hybridogenetic” vertebrate taxons: new
proposals. Alpes, 8: 61-14.

HaëckeL, E., 1868. — Natürliche Schôpfungsgeschichte. Berlin, Reimer: i-xvi + 1-568 + 204 b-c,

pl. 1-8+1.
HENNIG, W, 1950. — Grundzüge einer Theorie der phylogenetischen Systematik. Berlin, Deutscher Zen-
tralverlag: 1-370.

Mayr, E., 1974. - Cladistic analysis or cladistic classification? Z. 2001. Syst. Evol.-Forsch., 12: 94-128.

Le 1981. — Biological classification: toward a synthesis of opposing methodologies. Science, 214:
510-516.

—— 1982. - The growth of biological thought. Diversity. evolution, and inheritance. Cambridge, Mass. &
London, Belknap Press of Harvard Univ. Press: i-xiii

es 1995. — Systems of ordering data. Biol. & Phil, 10 (

MAYR, E. & ASHLOCK, P, 1991. — Principles of systematic zoology. Second edition. New York, McGraw-
Hill: i-xx + 1-475.

A

JBL. OÙ |
SÉUM)
PARS /

+ / Source : MNHN, Paris
/

AIVTES

International Journal of Batrachology
published by ISSCA

EDITORIAL BOARD

Chief Editor: Alain Dunois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,
25 rue Cuvier, 75005 Paris, France).

Deputy Editor: Janalee P. CaLDWELL (Oklahoma Museum of Natural History, University of Oklahoma,
Norman, Oklahoma 73019, USA).

Editorial Board Jean-Louis ALBARET (Paris, France); Ronald G. ALTiG (Mississippi State University,
USA); Francesco ANDREONE (Torino, Italy): Günter GOLLMANN (Wien, Austria): Tim HALLIDAY (Milton
Keynes, United Kingdom); W. Ronald Hever (Washington, USA); Masafumi MaTsut (Kyoto, Japan);
John C. Poynron (London, England); Ulrich Sinscu (Koblenz, Germany); Marvelee H. Wake (Berke-
ley, USA).

Technical Editorial Team (Paris, France): Alain Dupors (texts); Roger Bour (tables): Annemarie OHLER
(figures).

Index Editors: Annemarie OHLer (Paris, France); Stephen J. RICHARDS (Townsville, Australia).

SHORT GUIDE FOR AUTHORS
(detailed Instructions to Authors are given in Alytes, 1997, 14 (4): 175-200)

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with amphibians.
Beside articles and notes reporting results of original research, consideration is given for publication to synthetic
review articles, book reviews, comments and replies, and to papers based upon original high quality illustrations
(such as color or black and white photographs), showing beautiful or rare species, interesting behaviors, etc.

The title should be followed by the name(s) and address(es) of the author(s). The text should be
typewritten or printed double-spaced on one side of the paper. The manuscrit should be organized as follows:
Énglish abstract, introduction, material and methods, results, discussion, conclusion, French or Spanish
abstract, acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or tab. 4. Figures should not exceed
16 x 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures and tables
should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & POLLS PELAZ, 1989;
INGER et al., 1974). References in the Literature cited section should be presented as follows:

BourReT, R., 1942. - Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l’Indochine: i-x
+ 1-547, pl. I-IV.

Gr4r, J.-D. & POLLs PELAZ, M., 1989. - Evolutionary genetics of the Rana esculenta complex. In: R. M. DAWLEY
& J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The New York State
Museum: 289-302.

IxGer, R. F, Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some
Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet, 12: 121-145.

Manuscrits should be submitted in triplicate either to Alain Dupois (address above) if dealing with
amphibian morphology, systematics, biogeography, evolution, genetics or developmental biology, or to Janalce
P. CALDWELL (address above) if dealing with amphibian population genetics, ecology, ethology or life history.
Acceptance for publication wil be decided by the editors following review Dy at least (wo refetses

if possible aier acceptance à copy of the final manuscrpt on a floppy disk (3% r 5) should be sent
to the Chief Editor. We welcome the following formats of text processing: (1) preferably, MS Word (1.1 to 6.0,
DOS or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or WordStar (3.3 to 7.0); (2) less preferably,
formated DOS (ASCII) or DOS-formated MS Word for the Macintosh (on a 3 4 high density 1.44 Mo floppy
disk only).

Page charges are requested only from authors having institutional support for this purpose. The
publication of color photographs is charged. For each published paper, 25 free reprints are offered by ISSCA to
the author(s). Additional reprints may be purchased.

Published with the support of AALRAM
(Association des Amis du Laboratoire des Reptiles et Amphibiens
du Muséum National d'Histoire Naturelle, Paris, France).

Directeur de la Publication: Alain DuBois.
Numéro de Commission Paritaire: 64851.

©ISSCA 1997 Source : MNHN, Paris

Alytes, 1997, 15 (3): 105-136.

Contents

José P. POMBAL, Jr. & Cristiane A. MADUREIRA
A new species of Physalaemus (Anura, Leptodactylidae)
from the Atlantic rain forest of northeastern Brazil ................. 105-112

Abraham MUARES-URRUTIA
Un nuevo Leptodactylus (Anura, Leptodactylidae)
de un bosque nublado del oeste de Venezuela ....................... 113-120

Néstor G. Basso & Carmen A. UBEDA
The tadpole of Atelognathus nitoi

(Leptodactylidae, Telmatobiinae) .… 121-126
Indraneil Das & Romulus WHITAKER

A redescription of Ramanella mormorata Rao, 1937

(Anura MicrohVlidae) encens eee de ee re 127-132

Book review

Alain Dugois

An evolutionary biologist’s view

OnRE SCIENCE OR PIOIOS ET mn ee ee RU me ee 133-136

Alytes is printed on acid-free paper.
Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.
Dépôt légal: 4°" trimestre 1997.

© ISSCA 1997

Source : MNHN, Paris: